Paper · 2026 · Biology

Effects of Leaf Nutrients, Non-Structural Carbohydrates, and Microanatomical Structure on Biomass of Three Tree Species Under Drought Stress

Zhaoqun Ma; Xi Zhang; Mengyun Lei; et al.

20
claims
36
verbatim quotes
0
citations
2
research areas
✓
Open access · cc-by

Abstract

Drought stress profoundly affects plant growth and survival, but comparisons of integrated adaptive strategies across multiple tree species remain unclear. In this study, seedlings of Elaeagnus angustifolia (E. angustifolia), Populus euphratica (P. euphratica) and Xanthoceras sorbifolium (X. sorbifolium) were subjected to well-watered (CK), mild (T1), moderate (T2), and severe (T3) drought treatments. Leaf microanatomical traits, non-structural carbohydrates (NSCs), stoichiometric elements, biomass allocation, and key stress indicators were measured. The results showed that P. euphratica seedlings thickened leaves and vascular tissues and accumulated soluble sugars (SSs) and starch (ST) under T1–T2, but under T3, they prioritized root investment (root biomass +26.0%); their antioxidant enzymes were activated only under mild-to-moderate stress and declined under severe stress. E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass, continuously elevated activities of superoxide dismutase (SOD) and peroxidase (POD) as well as osmoregulatory substances (soluble protein SP, proline Pro), and showed the lowest malondialdehyde (MDA) content; their leaf carbon (C), nitrogen (N), and phosphorus (P) contents decreased the least, and their stoichiometric ratios remained stable. In contrast, X. sorbifolium seedlings progressively reduced leaf thickness and vascular area, depleted NSC reserves, exhibited unstable antioxidant responses, showed a significant decrease in Pro under severe drought, accumulated the highest MDA, and had the lowest N/P ratio, indicating the strongest nitrogen limitation. These results demonstrate that E. angustifolia combines structural plasticity, efficient nutrient use, robust osmotic adjustment, and sustained antioxidant capacity, conferring the strongest drought tolerance; P. euphratica* shows moderate tolerance through transient structural and carbon investment but suffers under extreme drought; X. sorbifolium has the weakest drought tolerance.

Abstract from the paper’s OpenAlex record; open copy licensed cc-by. Read at the source ↗

The text the model read, with the source record’s inline markup (italics, sub- and superscripts) rendered and its character entities decoded; quote offsets refer to the raw text.

Claims 20

Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Populus euphratica thickened leaves and vascular tissues under mild and moderate drought (T1-T2) compared with well-watered controls.

Taxon
P. euphraticaPopulus euphraticae1
Treatment
T1–T2e1
Outcome
leaves and vascular tissuese1
Direction
thickenede1
Comparator
well-watered (CK)e0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Populus euphratica
Populus euphratica · TaxonYoshi Canopus · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“subjected to well-watered (CK), mild (T1), moderate (T2), and severe (T3) drought treatments”

✓ located verbatim in the source · abstract, characters 311–403 · show in text ↑
e1

“P. euphratica seedlings thickened leaves and vascular tissues and accumulated soluble sugars (SSs) and starch (ST) under T1–T2, but under T3, they prioritized root investment (root biomass +26.0%)”

✓ located verbatim in the source · abstract, characters 581–777 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Populus euphratica accumulated soluble sugars and starch under mild and moderate drought (T1-T2) compared with well-watered controls.

Taxon
P. euphraticaPopulus euphraticae1
Treatment
T1–T2e1
Outcome
soluble sugars (SSs) and starch (ST)e1
Direction
accumulatede1
Comparator
well-watered (CK)e0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Populus euphratica
Populus euphratica · TaxonYoshi Canopus · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“subjected to well-watered (CK), mild (T1), moderate (T2), and severe (T3) drought treatments”

✓ located verbatim in the source · abstract, characters 311–403 · show in text ↑
e1

“P. euphratica seedlings thickened leaves and vascular tissues and accumulated soluble sugars (SSs) and starch (ST) under T1–T2, but under T3, they prioritized root investment (root biomass +26.0%)”

✓ located verbatim in the source · abstract, characters 581–777 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Populus euphratica increased root biomass by 26.0% under severe drought (T3), prioritizing root investment.

Taxon
P. euphraticaPopulus euphraticae1
Treatment
T3e1
Outcome
root biomasse1
Direction
increaseinferrede1
Value
+26.0%e1
Comparator
well-watered (CK)e0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Populus euphratica
Populus euphratica · TaxonYoshi Canopus · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“subjected to well-watered (CK), mild (T1), moderate (T2), and severe (T3) drought treatments”

✓ located verbatim in the source · abstract, characters 311–403 · show in text ↑
e1

“P. euphratica seedlings thickened leaves and vascular tissues and accumulated soluble sugars (SSs) and starch (ST) under T1–T2, but under T3, they prioritized root investment (root biomass +26.0%)”

✓ located verbatim in the source · abstract, characters 581–777 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Populus euphratica antioxidant enzymes were activated under mild-to-moderate stress.

Taxon
P. euphraticaPopulus euphraticae0
Treatment
mild-to-moderate stresse1
Outcome
antioxidant enzymese1
Direction
activatede1

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Populus euphratica
Populus euphratica · TaxonYoshi Canopus · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“P. euphratica seedlings thickened leaves and vascular tissues and accumulated soluble sugars (SSs) and starch (ST) under T1–T2, but under T3, they prioritized root investment (root biomass +26.0%)”

✓ located verbatim in the source · abstract, characters 581–777 · show in text ↑
e1

“their antioxidant enzymes were activated only under mild-to-moderate stress and declined under severe stress”

✓ located verbatim in the source · abstract, characters 779–887 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Populus euphratica antioxidant enzymes declined under severe stress.

Taxon
P. euphraticaPopulus euphraticae0
Treatment
severe stresse1
Outcome
antioxidant enzymese1
Direction
declinede1

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Populus euphratica
Populus euphratica · TaxonYoshi Canopus · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“P. euphratica seedlings thickened leaves and vascular tissues and accumulated soluble sugars (SSs) and starch (ST) under T1–T2, but under T3, they prioritized root investment (root biomass +26.0%)”

✓ located verbatim in the source · abstract, characters 581–777 · show in text ↑
e1

“their antioxidant enzymes were activated only under mild-to-moderate stress and declined under severe stress”

✓ located verbatim in the source · abstract, characters 779–887 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Elaeagnus angustifolia increased root biomass by 97.2% under severe drought (T3), with moderate leaf structural thickening.

Taxon
E. angustifoliaElaeagnus angustifoliae1
Treatment
T3e1
Outcome
root biomasse1
Direction
increasede1
Value
+97.2%e1
Qualifier
moderate leaf structural thickeninge1
Comparator
well-watered (CK)e0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Elaeagnus angustifolia
Elaeagnus angustifolia · TaxonGeorg Slickers · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“subjected to well-watered (CK), mild (T1), moderate (T2), and severe (T3) drought treatments”

✓ located verbatim in the source · abstract, characters 311–403 · show in text ↑
e1

“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”

✓ located verbatim in the source · abstract, characters 889–1041 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Elaeagnus angustifolia maintained stem biomass under severe drought (T3).

Taxon
E. angustifoliaElaeagnus angustifoliae0
Treatment
T3e0
Outcome
stem biomasse0
Direction
no_changeinferrede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Elaeagnus angustifolia
Elaeagnus angustifolia · TaxonGeorg Slickers · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”

✓ located verbatim in the source · abstract, characters 889–1041 · show in text ↑
Observationobservedoriginal resultpositivemeasured

Claim · model-written summary

Elaeagnus angustifolia showed continuously elevated superoxide dismutase and peroxidase activities.

Taxon
E. angustifoliaElaeagnus angustifoliae0
Outcome
activities of superoxide dismutase (SOD) and peroxidase (POD)e1
Direction
elevatede1
Qualifier
continuouslye1

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Elaeagnus angustifolia
Elaeagnus angustifolia · TaxonGeorg Slickers · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”

✓ located verbatim in the source · abstract, characters 889–1041 · show in text ↑
e1

“continuously elevated activities of superoxide dismutase (SOD) and peroxidase (POD)”

✓ located verbatim in the source · abstract, characters 1043–1126 · show in text ↑
Observationobservedoriginal resultpositivemeasured

Claim · model-written summary

Elaeagnus angustifolia showed continuously elevated soluble protein and proline.

Taxon
E. angustifoliaElaeagnus angustifoliae0
Outcome
soluble protein SP, proline Proe2
Direction
elevatede1
Qualifier
continuouslye1

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Elaeagnus angustifolia
Elaeagnus angustifolia · TaxonGeorg Slickers · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”

✓ located verbatim in the source · abstract, characters 889–1041 · show in text ↑
e1

“continuously elevated activities of superoxide dismutase (SOD) and peroxidase (POD)”

✓ located verbatim in the source · abstract, characters 1043–1126 · show in text ↑
e2

“as well as osmoregulatory substances (soluble protein SP, proline Pro)”

✓ located verbatim in the source · abstract, characters 1127–1197 · show in text ↑
Observationobservedoriginal resultpositivemeasured

Claim · model-written summary

Elaeagnus angustifolia had the lowest malondialdehyde content.

Taxon
E. angustifoliaElaeagnus angustifoliae0
Outcome
malondialdehyde (MDA) contente1
Direction
loweste1

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Elaeagnus angustifolia
Elaeagnus angustifolia · TaxonGeorg Slickers · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”

✓ located verbatim in the source · abstract, characters 889–1041 · show in text ↑
e1

“showed the lowest malondialdehyde (MDA) content”

✓ located verbatim in the source · abstract, characters 1203–1250 · show in text ↑
Observationobservedoriginal resultpositivemeasured

Claim · model-written summary

Elaeagnus angustifolia leaf carbon, nitrogen and phosphorus contents decreased the least under drought.

Taxon
E. angustifoliaElaeagnus angustifoliae0
Outcome
leaf carbon (C), nitrogen (N), and phosphorus (P) contentse1
Direction
decreased the leaste1

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Elaeagnus angustifolia
Elaeagnus angustifolia · TaxonGeorg Slickers · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”

✓ located verbatim in the source · abstract, characters 889–1041 · show in text ↑
e1

“their leaf carbon (C), nitrogen (N), and phosphorus (P) contents decreased the least”

✓ located verbatim in the source · abstract, characters 1252–1336 · show in text ↑
Observationobservedoriginal resultpositivemeasured

Claim · model-written summary

Elaeagnus angustifolia stoichiometric ratios remained stable under drought.

Taxon
E. angustifoliaElaeagnus angustifoliae0
Outcome
stoichiometric ratiose1
Direction
remained stablee1

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Elaeagnus angustifolia
Elaeagnus angustifolia · TaxonGeorg Slickers · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”

✓ located verbatim in the source · abstract, characters 889–1041 · show in text ↑
e1

“their stoichiometric ratios remained stable”

✓ located verbatim in the source · abstract, characters 1342–1385 · show in text ↑
Observationobservedoriginal resultnegativemeasured

Claim · model-written summary

Xanthoceras sorbifolium progressively reduced leaf thickness and vascular area under drought.

Taxon
X. sorbifoliumXanthoceras sorbifoliume0
Outcome
leaf thickness and vascular areae0
Direction
progressively reducede0

Verbatim

e0

“X. sorbifolium seedlings progressively reduced leaf thickness and vascular area”

✓ located verbatim in the source · abstract, characters 1400–1479 · show in text ↑
Observationobservedoriginal resultnegativemeasured

Claim · model-written summary

Xanthoceras sorbifolium depleted nonstructural carbohydrate reserves under drought.

Taxon
X. sorbifoliumXanthoceras sorbifoliume0
Outcome
NSC reservese1
Direction
depletede1

Verbatim

e0

“X. sorbifolium seedlings progressively reduced leaf thickness and vascular area”

✓ located verbatim in the source · abstract, characters 1400–1479 · show in text ↑
e1

“depleted NSC reserves”

✓ located verbatim in the source · abstract, characters 1481–1502 · show in text ↑
Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

Xanthoceras sorbifolium showed a significant decrease in proline under severe drought.

Taxon
X. sorbifoliumXanthoceras sorbifoliume0
Treatment
severe droughte1
Outcome
Proe1
Direction
decreasee1

Not reportedComparator

Verbatim

e0

“X. sorbifolium seedlings progressively reduced leaf thickness and vascular area”

✓ located verbatim in the source · abstract, characters 1400–1479 · show in text ↑
e1

“showed a significant decrease in Pro under severe drought”

✓ located verbatim in the source · abstract, characters 1546–1603 · show in text ↑
Observationobservedoriginal resultnegativemeasured

Claim · model-written summary

Xanthoceras sorbifolium accumulated the highest malondialdehyde content.

Taxon
X. sorbifoliumXanthoceras sorbifoliume0
Outcome
MDAe1
Direction
higheste1

Verbatim

e0

“X. sorbifolium seedlings progressively reduced leaf thickness and vascular area”

✓ located verbatim in the source · abstract, characters 1400–1479 · show in text ↑
e1

“accumulated the highest MDA”

✓ located verbatim in the source · abstract, characters 1605–1632 · show in text ↑
Observationobservedoriginal resultnegativemeasured

Claim · model-written summary

Xanthoceras sorbifolium had the lowest N/P ratio, indicating the strongest nitrogen limitation.

Taxon
X. sorbifoliumXanthoceras sorbifoliume0
Outcome
N/P ratioe1
Direction
loweste1
Qualifier
indicating the strongest nitrogen limitatione1

Verbatim

e0

“X. sorbifolium seedlings progressively reduced leaf thickness and vascular area”

✓ located verbatim in the source · abstract, characters 1400–1479 · show in text ↑
e1

“had the lowest N/P ratio, indicating the strongest nitrogen limitation”

✓ located verbatim in the source · abstract, characters 1638–1708 · show in text ↑
Observationauthor interpretationoriginal resultpositivebasis n/a

Claim · model-written summary

Elaeagnus angustifolia has the strongest drought tolerance of the three species.

Taxon
E. angustifoliaElaeagnus angustifoliae0
Outcome
drought tolerancee0
Direction
strongeste0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Elaeagnus angustifolia
Elaeagnus angustifolia · TaxonGeorg Slickers · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“E. angustifolia combines structural plasticity, efficient nutrient use, robust osmotic adjustment, and sustained antioxidant capacity, conferring the strongest drought tolerance”

✓ located verbatim in the source · abstract, characters 1741–1918 · show in text ↑
Observationauthor interpretationoriginal resultmixedbasis n/a

Claim · model-written summary

Populus euphratica shows moderate drought tolerance but suffers under extreme drought.

Taxon
P. euphraticaPopulus euphraticae0
Outcome
tolerancee0
Direction
moderatee0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Populus euphratica
Populus euphratica · TaxonYoshi Canopus · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“P. euphratica* shows moderate tolerance through transient structural and carbon investment but suffers under extreme drought”

✓ located verbatim in the source · abstract, characters 1920–2044 · show in text ↑
Observationauthor interpretationoriginal resultnegativebasis n/a

Claim · model-written summary

Xanthoceras sorbifolium has the weakest drought tolerance of the three species.

Taxon
X. sorbifoliumXanthoceras sorbifoliume0
Outcome
drought tolerancee0
Direction
weakeste0

Verbatim

e0

“X. sorbifolium has the weakest drought tolerance”

✓ located verbatim in the source · abstract, characters 2046–2094 · show in text ↑